wood decay fungi in hornbill nest cavities in khao yai...

95

THE RAFFLES BULLETIN OF ZOOLOGY 2011

WOOD DECAY FUNGI IN HORNBILL NEST CAVITIES IN KHAO YAI NATIONAL PARK, THAILAND

Sirirak Supa-AmornkulDepartment of Microbiology, Faculty of Science, Mahidol University, Bangkok, Thailand

Suthep WiyakruttaDepartment of Microbiology, Faculty of Science, Mahidol University, Bangkok, Thailand

Pilai Poonswad*Department of Microbiology, Faculty of Science, Mahidol University, Bangkok, Thailand

E-mail:[email protected] (*Corresponding author)

ABSTRACT. – We investigated the abundance and species diversity of wood decay fungi inside and outside cavities of predominant hornbill nest trees, Dipterocarpus gracilis and Cleistocalyx nervosum and compared with those of Balakata baccata, which is known as non-nest tree for lack of cavity, in a moist evergreen forest of Khao Yai National Park, Thailand. We also examined the abundance of cavities in these tree sampled from wild population. Wood samples were collected twice yearly, once in dry and the other in wet season, from inside and outside the cavities of 10 D. gracilis trees, 10 C. nervosum trees, and 6 B. baccata trees. Additional dead wood of Dipterocarpus and Cleistocalyx was collected in dry season to determine true wood decay fungi. A total of 1,199 fungal isolates were obtained and were classifi ed to 68 species, 52 genera, 33 families, and 4 phyla The number of isolates was signifi cantly higher from inside than outside the cavities and in wet season than in dry season, particularly the isolates from Dipterocarpus indicating wet season provides optimal condition for fungi. Cleistocalyx hosted the highest species richness, i.e. 51 species (75%). Forty species (58.8%) were identifi ed from Dipterocarpus, and as expected the lowest was from Balakata, with only 16 species (23.5%). Eight species (11.8%) were found in common among all trees studied while 10, 20 and 4 species were exclusively isolated from the respective trees. Simpson’s and Shannon-Weiner indices were used to determine fungal species diversity. In dry season, Dipterocarpus hosted the highest diversity according to Simpson’s index, while Cleistocalyx had the lowest diversity, as demonstrated by both indices, in fungal species. Unexpectedly, wood decay fungi were dominated by soft rot fungi (91.2%) but not white rot (5.9%). The most common soft rot fungi were Trichoderma sp., Gliocladium sp. and Fusarium sp., and the white rot were Coprinus sp. and Sporotrichum sp. The present study suggested that wet season and enclosure condition such as inside the cavity infl uence the abundance and species richness. Other than moisture, additional conditions inside the cavity that may enhance the abundance and species richness of fungi included the volume of substrate (dead wood) and remaining nest debris (fruits, insects, and old feathers of imprisoned female). Further study on nature of sap and chemical compounds produced by Balakata is needed in order to explain the lack of large size cavity in Balakata, despite being a softwood tree. Nest cavities of hornbills were predominantly found in two tree genera/species, Yang sian (Dipterocarpus gracilis) and Wa (Cleistocalyx nervosum). By contrast, Sali nok (Balakata baccata), although it is large in size and common in the study area, no nest cavity was found. In this study, the abundance of cavities in these three tree genera was determined. The numbers of trees with at least one cavity were the lowest for Balakata (4%), 15.5% for Dipterocarpus, and 13.7% for Cleistocalyx. The trends were also similar for the abundance of cavities. The abundance of cavities in these tree genera was somewhat related to susceptibility of the tree to wood decay fungi. The susceptibility (or resistance) and host specifi city may be related to wood structure and/or chemical substance the tree produced, which may inhibit (or enhance) growth rate of fungi and in turn affect decay process.

This paper was presented at the 5th International Hornbill Conference jointly organised by the National Parks Board (Singapore) and the Hornbill Research Foundation (Thailand), in Singapore on 22nd–25th March 2009.

KEY WORDS. – Wood decay fungi, hornbill nest cavity, Dipterocarpus gracilis, Clesitocalyx nervosum, Balakata baccata, Khao Yai National Park.

THE RAFFLES BULLETIN OF ZOOLOGY 2011 Supplement No. 24: 95–113Date of Publication: 30 Mar.2011© National University of Singapore

INTRODUCTION

Wood decay fungi are important to ecological systems as they are responsible for recycling of plant materials and

are important components of global carbon cycling (Odor et al., 2006). Their ability to degrade wood components is crucial to wood decomposition process. Three groups of wood decay fungi are recognized: white rot, brown rot and

96

Supa-Amornkul et al.: Wood decay in hornbill nests

soft rot fungi based on the components of wood that they can degrade (Käärik, 1974). Different groups have different abilities in degrading different substances. For instance, the white rot Basidiomycetes and xylariceous Ascomycetes are the principal organisms responsible for wood decay in hardwood of tropical forests. In contrast, brown rot fungi are important in coniferous forest (Hammel, 1997). Wood decay fungi utilize a variety of starting substrates, including living trees with heart and butt rot, snags or fallen logs (Lonsdale et al., 2008). Wood decay fungi thus play an important role in creating ecological niches (Lonsdale et al., 2008), including cavities in trees, and at the same time degrading dead wood of forest trees in a forest ecosystem.

How do wood decay fungi get involved in biology of hornbills, particularly in breeding biology? Hornbills are very large birds of the family Bucerotidae that inhabit tropical forest and savanna of Asia and Africa (Kemp, 1995) and are secondary-cavity nesters. They are unable to excavate their own nest cavity, and therefore strictly rely on a suitable existing cavity, which is recognized as an important breeding limiting factor (Poonswad, 1995). The role of wood decay fungi as dead wood decomposers in the forest ecosystem indirectly benefi ts hornbills by creating and/or enlarging a cavity, which may later serve as a nest chamber.

The availability of cavities is very important to a wide variety of animals ranging from insects, reptiles, birds to mammals, who utilize them as nesting chambers or shelters (Poonswad, 1993). In a tropical forest, a cavity in a tree can be created by a number of ways, directly or indirectly. A cavity can be actively excavated by animals, particularly birds such as woodpeckers (Family Picidae) (Hopper & Lennartz, 1991), with an assumption that eventually the cavity becomes infected by wood decay fungi and gets enlarged. The cavity can also be indirectly created in trees with physical damage caused by natural phenomena, frequently by wind. The wind breaks off branches and causes wounds on the tree trunk (Poonswad, 1995), which subsequently is subjected to infection by one or more wood-degrading microorganisms, including wood decay fungi. In a tropical forest, such as moist evergreen forest, the study of wood decay fungi is very limited (Lodge, 1997). There is no documentation on species richness inside cavities of tropical forest trees, and whether the species richness has correlation with the existence and the size of the cavity. Known information on wood decay fungi that infect tropical trees of family Dipterocarpaceae and cause heart and butt rot are fungi of genera Ganoderma and Cryptoderma (Chalermponse, 1987); and presumably a cavity is formed. Trees in family Dipterocarpaceae i.e., Dipterocarpus, Hopea, Shorea, Neobalanocarpus and Syzygium of Myrtaceae are known to be predominant trees, which bear nest cavities for hornbills in Thailand, (Chuailua et al., 1998, Poonswad et al., 2005). Wood of various species of Syzygium, on the other hand, demonstrated resistance to white and brown rot fungi tested for decay resistance in Fiji (Osboren & Lynette, 1967). Wood decay fungi do not involve only in creating cavities but also degrading cavities. This obviously has a negative effect on the hornbill. The continuation of decay process inevitably causes the cavity fl oor to sink deeply,

thus making the cavity unsuitable as a nest for hornbills. Such cause is recognized as the main problem found in the nest trees of Dipterocarpus and Syzygium genera (Chuailua et al., 1998). The loss of suitability of a nest cavity directly affects the reproduction of hornbills and hornbill population as a whole.

In this paper, the fi rst study in a moist evergreen forest, we investigated abundance and diversity of wood decay fungi both inside and outside cavities of hornbill nest trees. We also extended our study to investigate the wood decay fungi on non-nest trees in this forest type.

MATERIALS AND METHODS

Study area. – The study was carried out in a moist evergreen forest, the habitat of hornbills, at Khao Yai National Park situated in the central-north region of Thailand (14° 15' to 14° 35'N and 101° 5' to 101° 52'E) (Fig. 1). In the park, there are four hornbill species living in sympatry, viz. Great Hornbill (Buceros bicornis), Wreathed Hornbill (Rhyticeros undulatus), White-throated Brown Hornbill (Ptilolaemus austeni) and Oriental Pied Hornbill (Anthracoceros albirostris) (Poonswad et al., 2005). Within the study area, dominant and common large trees (diameter at breast height or dbh ≥ 40 cm) are Dipterocarpus gracilis (referred to as D. costatus in Liewviriyakit, 1989), Sloanea sigun, Castanopsis accuminatissima, Cliestocalyx nervosum (referred to as Eugenia sp. in Liewviriyakit, 1989), Choerospondias axillaris, and Balakata baccata (referred to as Sapium baccatum in Poonswad, 1993).

Collection of wood samples. – Wood samples were collected from cavities that were used by hornbills but abandoned during the study. These cavities situated in trees known as predominant nest tree species i.e., Dipterocarpus gracilis (Dipterocarpaceae) (referred to as D. costatus in Liewviriyakit, 1989) and Cleistocalyx nervosum (Myrtaceae) (referred to as Eugenia cumini in Liewviriyakit, 1989 and Eugenia sp. in Poonswad, 1995, and Syzygium sp. in Chuailua et al., 1998, Poonswad, 1995). Twenty hornbill nest cavities, ten in D. gracilis (Dipterocarpus hereafter, otherwise stated) and ten in C. nervosum (Cleistocalyx hereafter, otherwise stated), that were accessible were selected from abandoned nests with unknown cause and regardless of previous hornbill species. The mean height above ground of these cavities was 25.1±6.0 m in Dipterocarpus and 15.9±3.1 m in Cleistocalyx. To access these hornbill nest cavities, rope climbing was used. For control, wood samples were collected from six Balakata baccata trees (Euphorbiaceae) (Balakata hereafter, and referred to as Sapium baccatum in Poonswad, 1993), a non-hornbill nest tree species (Poonswad, 1993) with dbh ≥ 40 cm (the smallest size of tree which bears hornbill’s nest, Poonswad, 1995), a common large tree species (Gardner & Sidisunthorn, 2000) at about 60 cm above ground, and an existing scar or wound on the tree trunk. Locations of studied trees were marked with GPS (Garmin, eTrex) and are shown in Fig. 1. Wood samples from all trees were collected in dry season (between December 2005 and March 2006).

97


Fig. 1. Location of Khao Yai National Park and locations of sampled trees.

In wet season, we were unable to collect samples during consecutive months due to diffi culties in accessing the cavities of Dipterocarpus. Only wood samples from Cleistocalyx were collected (between June and July 2005). Wood samples from Dipterocarpus and Balakata were then collected between August and October, 2006. Additionally, wood samples from six dead Diptercarpus and six Cleistocalyx trees were collected in dry season (March 2006) for the observation of true wood rot fungi.

Wood samples were collected at various positions of the cavity using a chisel. Inside the nest cavity, samples were collected from left and right sidewalls and the wall opposite the cavity opening or nest entrance. Outside samples were collected at the backside opposite the cavity opening. From Balakata, samples were taken from a wound or a scar found on the trunk.

Abundance of cavities. – Abundance of tree cavities with dbh ≥ 40 cm from wild populations was checked. We sampled predominant nest-tree species from 103 Dipterocarpus trees, 102 Cleistocalyx trees, and 101 non-nest Balakata trees along Thanarat Road inside the park between Km 31-34 posts. The number of cavities found was recorded regardless of the cavity size or position of the cavity on the tree. Measurement of individual tree size (in dbh) was not done.

Fungal isolation. – Lignin cellulose medium (LCM) was modifi ed from minimal medium (Paterson & Bridge, 1994) by adding lignin cellulose as a carbon source instead of sucrose. LCM was modifi ed to facilitate the growth of fungi which can degrade wood components. The LCM agar was used to select wood decay fungi from wood samples. Half potato dextrose agar (1/2 PDA) was used to purify the fungal isolates. For identifi cation of the fungi, malt extract agar (MEA), potato dextrose agar (PDA) (Merck), sabouraud 4% dextrose agar (Merck) (SDA) and sporulated medium agar (SA) (Nugent et al., 2006) were used to determine macroscopic morphology of the fungi. When molecular identifi cation was necessary, potato dextrose broth (PDB) was used to prepare fungal mycelium.

Fungi identifi cation. – Fungal isolates were classifi ed to genus and species when possible. For sporulating fungi, we identifi ed morphological characteristics through observations, both macroscopically and microscopically. For uncertain and non-sporulating fungi, molecular technique was used. Briefl y, Whatman FTA card was used for extraction of fungal DNA of non-sporulating fungi. ITS1-5.8S-ITS2 regions of ribosomal DNA (rDNA) used for fungal identifi cation were amplifi ed by PCR using the forward primer ITS1 and the reverse primer ITS4 as described in White et al., (1990). Amplifi cation reaction was performed in a total volume of 50 µl. Reaction mixture contained 0.5 µM of each primer, 50 µM of individual dNTP, 3 mM of MgCl2, 10X buffer, 1 U Dynazyme EXT polymerase (Finnzyme) and a processed 2-mm FTA dish carrying the fungal DNA. Amplifi cation process consisted of (i) pre-denaturation step at 95°C for 3 min, (ii) 30 consecutive cycles at 95°C for 50 sec (denaturation), 55°C for 40 sec (annealing) and 72°C for 40 sec (extension), and (iii) a fi nal extension at 72°C for 10 min. PCR products were ligated into pGEM-T easy vectors for preparation of recombinant plasmids and transformed into E. coli DH5α. Isolated plasmid from each clone was checked for insertion by restriction enzyme digestion. Extracted plasmids were sequenced in an automate sequencer (ABI PRISM TM model 3730XL, Perkin Elmer). The fungi were identifi ed by comparing the ITS1-5.8S-ITS2 sequences with those of the reference strain fungi in the GenBank.

Data analyses. – To compare the abundance of cavities found in Dipterocarpus, Cleistocalyx and Balakata trees, Chi square-test was used. The fungal species richness isolated from each studied tree species was compared by rarefraction method. To measure diversity of isolated fungi, the Simpson’s reciprocal and Shannon-Wiener indices were used. To test the correlation between the numbers of isolates and species, Spearman Rank Correlation was used. To determine similarity of isolated fungi in various host trees, Chao-Sorensen Est-Abundance Base was applied (Chao et al., 2005). Species richness and diversity indices were calculated by BioEdit version 7.0. All statistic tests were calculated by SPSS version 13.00 program.

RESULTS

Abundance of cavities in studied trees. – The numbers of cavities found in wild tree population were relatively low, only 34 (11.1%) out of 306 trees (Table 1). There was no difference in the numbers of cavities recorded from Dipterocarpus and from Cleistocalyx trees, (χ2=0.668, df=1, P=0.414) (Table 1). Balakata, as non-nest trees, obviously had a low number of cavities. Therefore, Balakata had signifi cantly less cavities than those of the former two species (χ2=9.78, df=1, P=0.02 and χ2=6.093, df =1, P=0.014, respectively). The number of cavities per tree was highest in Cleistocalyx (1.8 cavities /tree) (Table 1). It was also noticed that cavities found in Balakata were shallow and small.

Fungal abundance and seasonality. – A total of 1,199 fungal isolates were obtained from studied trees and two additional

98


Table 1. Number of cavity trees and average number of cavities observed in sampled trees from wild populations of Dipterocarpus, Cleistocalyx, Balakata at Khao Yai National Park.

Tree species N No. cavity tree (%) Total cavity Ave. number of cavities per tree

Diptercarpus 103 16 (15.5) 17 1.1

Cleistocalyx 102 14 (13.7) 25 1.8

Balakata 101 4 (4.0) 4 1

Total 306 34 (11.1) 46

dead Dipterocarpus and Cleistocalyx (Table 2). Of the total, the number of isolates obtained from Dipterocarpus and Cleistocalyx in both wet and dry seasons were similarly high, 565 isolates (47.1%) and 513 (42.8%), respectively (Table 2). However, the highest number of isolates (429 isolates) was from the cavities of Dipterocarpus in wet season. To compare the total number of isolates from these nest trees and that from Balakata, we also included the isolates from outside the cavity. In wet season, the number of isolates was apparently higher than that in dry season for all studied trees (Table 2, Fig. 2a, and Appendices 1a–b). Distinct increase in the number of isolates in wet season indicated that season has high infl uence on the abundance of fungi, especially on Dipterocarpus and Cleistocalyx (Appendices 1a–b).

The majority of isolates, 1,182 (98.6%), sporulated in cultures, and 90.9% (1,074 isolates) were from Dipterocarpus and Cleistocalyx. The rest (17 isolates, 1.4%) were non-sporulating fungi and were mainly from Balakata in wet season (10 isolates, 58.8 %) (Table 2).

Fig. 2. a) Total number of fungal isolates from 10 of studied trees in different seasons and b) Total number of fungal species isolated from studied trees in different seasons.

To compare the average number of isolates from inside and outside the cavity, we consistently calculated the number of isolates from samples collected from one location, the right wall of the cavity, since the average numbers of isolates obtained from various locations inside the cavity showed no signifi cant difference. Therefore, the data shown in Table 3 and Fig. 2 were from the right wall inside the cavity. For the rest of the analyses, we used entire data from all samples collected inside and outside (Appendices 1a–b), otherwise stated. Fig. 2a compares total number of fungal isolates obtained from inside and outside the cavities of Dipterocarpus, Cleistocalyx and Balakata in dry and wet seasons. The average number of fungal isolates collected from inside and outside cavities among these studied trees in both seasons were signifi cantly different (Kruskal-Wallis test, χ2=54.756, df=3, P=0.000) (Table 3). The average number of isolates was the highest, both from inside and outside cavity of Dipterocarpus, in wet season and signifi cantly differed from those of Cleistocalyx. Overall, pairwise test shows signifi cant difference between the average numbers of isolates from inside and outside in both seasons, except for a few pairs (Table 3). It is apparent that wet season and location (i.e., inside the cavity) had infl uence on the abundance of fungi (Table 3 and Fig. 2a). The isolate abundance was found to be very highly correlated with the number of species (Spearman Rank Correlation: R2 0.900, P=0.000, N=52) (Fig. 2). Therefore, both terms, isolates and species, were used in the following analyses. Species richness had the same trend as the abundance of isolates i.e., high in wet season and from inside the cavities (Table 2, Fig. 2b, and Appendices 1a–b).

Species richness, diversity and composition. – The species accumulation curves for species richness of fungi isolated from different studied tree species shows that species added per tree appeared to reach or nearly reach stationary state for all host tree species (Fig. 3a). Similar trend, species accumulation curves obtained from outside the cavity or on the trunk of each studied trees nearly reached stationary state except for those of Balakata (Fig. 3b). The highest number of species was recorded from Cleistocalyx trees (Table 2, Fig. 3, and Appendix 1a). However, additional number of trees to each host tree/wood may yield a slightly more genera/species. Hence, number of sample size used in this study was considerably suffi cient.

List of fungal isolates and species recorded from inside and outside of all studied trees and wood in dry and wet seasons is given in Appendix 1. Of the 1,199 isolates, there were 68 species (43 unidentifi ed) of 52 genera from 30 families in 4 phyla identifi ed (Table 2 and Appendix 1a). Among the

99


Tab

le 2

. Sum

mar

y of

fun

gal

isol

ates

ide

ntifi

ed i

n th

is s

tudy

. Sho

wn

here

are

the

num

bers

of

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d no

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num

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nd

dive

rsity

ind

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and

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for

each

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D

ry

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B

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D

ip

Cle

is

BB

D

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Dip

D

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Cle

is

Dip

C

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B

B

Tota

l D

ip

Cle

is

BB

(n

=10)

(n

=10)

(n

=6)

(n=6

) (n

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(n=1

0)

(n=1

0)

(n=6

)

(n=2

0)

(n=2

0)

(n=1

2)To

tal

(Iso

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136

195

21

22

37

429

318

41

1199

56

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3 62

Tota

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a/Sp

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s)

23

27

9 11

10

33

40

12

68

40

51

16

Sp

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Sp

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s 23

27

9

10

10

32

37

10

62

Gen

era

16

20

8 7

4 22

26

7

45

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9 11

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14

19

8 28

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6 19

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42

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11

82

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100


Fig. 3. a) Species accumulation curves obtained from each type of studied tree species/wood and b) species accumulation curves obtained from outside-cavity or tree- trunk isolates for each type of studied tree species/wood.

68 species, 40 species (58.8% of total) were identifi ed from Dipterocarpus, 51 (73.5%) from Cleistocalyx, 16 (23.5%) from Balakata, 11 (16.2%) from dead Dipterocarpus and 10 (14.7%) from dead Cleistocalyx. There were 10 exclusive species (14.7%) found in Dipterocarpus, 20 (29.4%) in Cleistocalyx, four species (5.9%) in Balakata, and only one species (1.5%) i.e., Trichoderma spirale in dead Dipterocarpus. There were eight species (11.8%) found in common among these three studied species (Table 2 and Appendix 1a). Among 30 exclusive species found in both species of nest trees, there were 23 species (76.7%) found only inside the cavity and four species (13.3%) found outside the cavity regardless of host tree species (Appendix 1b).

There were three unidentified species of Fusarium, Gliocladium and Trichoderma genera found in most abundance in both seasons (Appendix 1a). Of 68 species obtained, 14 (20.6%) were isolated only once. Five isolates could not be identifi ed to species level (Appendix 1a).

Considering diversity indices given in Table 2, we compared between Dipterocarpus and Cleistocalyx based on the same data collection from both inside and outside the cavities. The highest diversity was from Dipterocarpus in dry season determined by both Simpson’s and Shannon-Weiner indices (10.30 and 1.14, respectively). In contrast, fungal diversity in Cleistocalyx was markedly high in wet season, (9.02 and 1.21, respectively) (Table 2). To compare species diversity index among 3 studied tree species, we used data from outside the cavity. Dipterocarpus hosted highest species diversity of

wood decay fungi in dry season (Simpson’s index: 11.34), but the diversity markedly decreased in wet season based on Simpson’s index. In contrast, species diversity in Cleistocalyx was the lowest in dry season, indicated by both Simpson’s and Shannon-Weiner indices (3.33 and 0.44, respectively) and increased drastically (6.30 and 0.91, respectively) in wet season. On the other hand, species diversity in Balakata was similar in both seasons. It should also be noted that the dead Cleistocalyx hosted low fungal diversity as confi rmed by both values of Simpson’s and Shannon-Weiner indices (4.47 and 0.77, respectively).

The ten most common fungal species ranked by the number of isolates from each studied tree genus and dead wood are shown in Table 4 and accounted for a total of 25 species from fi ve different sample types (trees and wood). Scoring was obtained from ranking of fungal species by host trees. The rank of each fungal species from each tree/wood was scored ranging from 10 to 1 for corresponding rank 1 to 10, then combined to get fi nal rank for each fungal species. Trichoderma sp. 1 was the most frequently isolated in this study, followed by Gliocladium sp. 1 and Fusarium sp. 1 (Table 4).

Wood decay fungi found in this study were dominated by soft rot fungi (62 species, 91.2% of the total 68) (Table 5). Among these soft rot fungi, 58 of 62 species (93.5%) were in Ascomycota (Appendix 1a). There were only two species of white rot fungi i.e., Coprinus sp. isolated only from outside of Dipterocarpus, and Sporotrichum sp. isolated from both inside and outside cavities of both Dipterocarpus and Cleistocalyx (Table 5 and Appendix 1a). Soft rot fungi were isolated in all studied trees with similar percentages (Table 5). While white rot fungi were isolated from cavities of Dipterocarpus and Cleistocalyx, but not from Balakata. Candelariella sp., Candida sp., Carpoligana sp., and Phaetrichospaeria sp. were not wood decay fungi (Table 5 and Appendix 1).

DISCUSSION

Fungal abundance and species richness. – Among limited studies on wood decay fungi in tropical forest, most of wood documented decomposition by wood decay fungi (Lonsdale et al., 2008; Osboren & Lynette, 1967; Setala & MacLean, 2004) and host specifi city (Gilbert & Sousa, 2002, May, 1991), but not the species richness of wood decay fungi from tree cavities. More numbers of fungal isolates and species richness in wet season than in dry season for all sample trees in this study (Tables 2–3; Fig. 2a–b; Appendix 1) indicate the infl uence of physical environment factors on the growth of fungi, which appears to be in agreement with Käärik (1974) who suggested that the most important factors in spreading and survivals of microorganisms in wood are moisture content of the wood, aeration, temperature and interaction between microorganisms. Being a suitable host for any microorganisms, there is one important process i.e., establishment. The establishment of wood decay fungi is known to dependent on combined factors, including infection source (e.g., arrival of spores), optimal physical

101


Tab

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f fu

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and

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Sign

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ank

test

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st

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χ2 (d

f=3)

P

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Z

pair

Dip

-2.7

10

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DO

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7 9

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.

102


Table 4. The ten most prevalent genera/species recorded from each studied tree and dead wood with sum scores and fi nal ranks.

Species

Rank Sum Score Final Rank

Dip Cleis BB DD DC

Trichoderma sp1. 2 1 1 1 1 49 1

Gliocladium sp1. 1 4 5 6 4 35 2

Fusarium sp1. 5 2 6 6 2 34 3

Gliocladium virens 3 3 6 21 4

Fusarium solani 8 8 3 6 19 5

Fusarium ventricosum 6 5 5 17 6

Trichoderma harzianum 2 3 17 6

Sporotrichum sp. 3 6 13 8

Paecilomyces victoriae 3 6 13 8

Aspergillus sp. 9 4 9 10

Bipolaris oryzae 2 9 10

Cylindrocarpon sp. 7 6 9 10

Mucor sp. 3 8 13

Penicillium sp. 3 8 13

Acremonium sp. 8 6 8 13

Phialophora richardsiae 6 5 16

Trichoderma spirale 6 5 16

Paecilomyces sp. 6 5 16

Pythium sp. 7 4 19

Mortierella sp. 7 4 19

Arthrographis cuboidea 8 3 21

Codinaea sp. 8 3 21

Carpoligna sp. 8 3 21

Verticillium sp. 10 1 24

Cylindrocladium sp. 10 1 24

The lowest value of the rank is genus/species which has the highest number of fungal isolates. Dead=dead Dipterocarpus and Cleistocalyx wood sample.

Table 5. Numbers of genera/species and percentages of each wood decay fungal group found in studied trees and dead wood.

Fungal group Dipterocarpus Cleistocalyx Balakata Dead Dip Dead Cleis Total No. % No. % No. % No. % No. %

Soft rot 36 90 48 94.1 15 93.8 11 100 10 100 62

White rot 2 5 1 2 0 0 0 0 0 0 4

Non rot 2 5 2 3.9 1 6.2 0 0 0 0 2

Total genera/species 40 51 16 11 10 68

Dead Dip=dead Dipterocarpus wood, Dead Cleis=dead CLeistocalyx wood.

environment conditions (e.g., temperature and moisture), suitable substrate, and absence of poisonous or inhibiting substances (Käärik, 1974). Not only external factors or physical factors are important, the establishment is certainly infl uenced by biological factors, for instance, plant cell components, which form wood structure and properties. Cell walls of these have different structures which resist to different fungal enzymes (Akin et al., 1995; Gilbert & Sousa, 2002) at different levels.

This study strongly suggests that an important external factor for growth of fungi is moisture. Fungal species richness in

the studied trees during wet season and inside the cavity (Tables 2–3; Fig. 2; Appendix 1) clearly favors moist condition. Inside the cavity, moisture may not differ much in both dry and wet seasons. Thus, species richness in both Dipterocarpus and Cleistocalyx was relatively similar, except for inside Cleistocalyx in wet season which accommodated the highest species richness (Fig. 2b; Appendix 1b). Relative humidity inside a hornbill nest cavity measured 24 hours varied between 70% and 100% and daily mean temperature varied between 22°C and 27°C in dry season and in the same area as the present study (Poonswad, 1993). These ranges of environmental factors are perhaps suitable for the

103


establishment and growth of the fungi. On the other hand, daily relative humidity on the ground varied greatly between 40% and 100% and mean temperature varied between 19°C and 23.3°C in dry season (Poonswad, 1993). This may not favor the growth of fungi outside the cavity as shown by pairwise test (Table 3). The number of fungal isolates from Balakata (only outside) did not differ in both seasons but was signifi cantly less than those of the other two species in both seasons. This moisture content does not correspond to the case of Balakata. If the decay rate is related to the ability of fungi to establish, which may further be implied to species richness, then the effect of wood moisture content on decay rate documented by Käärik (1974) would be sensible. As wood of Balakata contains very high moisture content (187%, Appendix 2); therefore, it may inhibit the establishment of the fungi. Besides high moisture content, this case suggests that there might be other factors which reduce the susceptibility or establishment. For instance, sap and chemical compounds produced by this tree may have defensive properties or activities which prevent further infection after an injury occurs (e.g., after sample collection in the fi rst season). Therefore, it is interesting to study fungi inside those small and shallow cavities of Balakata and nature of its sap, particularly chemical compounds produced and their activities.

Other than moisture and temperature, factors infl uencing species richness inside the cavity include physical environments and nest debris (e.g. fruits and insect remains). It is known that imprisoned female and chicks are fed with a great variety of fruits and insects by the male (Poonswad et al., 1998). These debris and remains provide optimal condition and additional substrates for fruit rot fungi. High species richness inside the cavities of Dipterocarpus and Cleistocalyx may be infl uenced by these fungi living on such debris inside the cavity. High number of isolates in Dipterocarpus was greatly contributed by the isolates of Gliocladium sp., which is one of the documented fruit rot fungi (Appendix 1a).

Wood decay fungi and cavities. – In this study, the initiation of a cavity in these studied trees is not our prime objective. For hornbills, as secondary cavity nesters, are unable to excavate their own nests (Poonswad et al., 1987). Cavities in trees speculated to be the results of interactions between wood of host tree (substrate) and decomposers, including wood decay fungi, after the trees have been injured by any causes. There are a number of studies suggesting that the rate of wood decomposition by the complex of wood-colonizing microorganisms, including wood decay fungi, tends to be affected by seasonal variations of temperature and moisture content in wood (Käärik, 1974; Lonsdale et al., 2008; Poonswad, 1995). Wood decay fungi play important roles in hornbill’s life through decomposition process: positive effect (e.g., creation of a cavity) and negative effect (e.g., degrading a nest chamber). Being predominant nest tree species, the Dipterocarpus and Cleistocalyx and the non-nest tree, the Balakata, were confi rmed by the corresponding number of cavities recorded in wild population and the abundance of wood decay fungi isolated (Tables 1, 2, 5, Fig. 2b and Appendix 1).

Study on the relationship between species richness and decomposition rate is inconclusive. However, there are studies on species richness of soil fungi that had positive relationship with litter decomposition rate (Setala & MacLean, 2004; Tiunov & Scheu, 2005). Käärik (1974) documented that wood with moisture content between 60 and 100 % would be rapidly decomposed, and those contains moisture lower than 30% or higher than 120% would not be putrefi ed. If consider only wood moisture content, only a few and small cavities would have been expected from Cleistocalyx and Dipterocarpus. While Cleistocalyx had highest number of cavities and highest species richness of wood decay fungi (Tables 1–2), wood of this species contains 12 % of moisture content, which is the same as Dipterocarpus (Appendix 2). We hypothesize that toughness and strength of wood affect fungi susceptibility and decay rate, as observed in the tree with tougher wood i.e., Dipterocarpus, which has reduced fungi susceptibility and decreased decay rate, when compared to Cleistocalyx (Appendix 2). On the contrary, Balakata wood properties (including toughness, strength and hardness) are far less than those of the former two species, except for the very high moisture content (187%). However, Balakata wood had the least number of cavities and the lowest species richness. Other evidence, which may better explain the difference in fungal species richness, by Lonsdale et al. (2008) stated that species richness of wood decay fungi positively related with the amount of substrate. Therefore, the substantial differences in fungal species richness between those of Cleistocalyx or Dipterocarpus and Balakata can be explained by the fact that the former two species had greater volume of substrates (dead wood) inside the cavity but lack in the latter species.

However, studies on wood decay fungi and their host trees suggest that structure of plant cells and antifungal components in each plant species infl uence decay process by wood decay fungi (Akin et al., 1995; Alfredsen et al., 2008; Gilbert & Sousa, 2002). Major components of wood cell wall are cellulose, hemicelluloses, and lignin. Wood decay fungi of different groups have different ability in breaking down these substrates (Käärik, 1974). Lacking information on wood cell wall components of all trees in this study, it is not possible to make a conclusion that wood components are involved as factors in wood degradation by wood decay fungi. The highest species richness of wood decay fungi isolated from inside cavities of Cleistocalyx trees in this study (Tables 2, 5, Fig. 2b and Appendix 1b) and higher percentage of nest cavities with sunken fl oor in Cleistocalyx than those in Dipterocarpus (Chuailua et al., 1998) may imply that the rate of wood decomposition inside the cavity of Cleistocalyx is higher than that of the Dipterocarpus trees, which hosted less species richness (Tables 2 and 5, Fig. 2b). As availability of suitable nests is important for hornbill population, wood decay results in relatively high percentage of cavity loss, which directly affects hornbill breeding. The sunken nest fl oor, as a result of decomposition process, in these two predominant nest trees may be enhanced by relative humidity and temperature inside the cavity (70–100% and 22–27°C, respectively, Poonswad, 1993), although the wood of these two species has low moisture content.

104


Wood decay fungi isolated in this study were dominated by soft rot fungi (62 species or 91.2% of total species, Table 5). This composition is similar to the fungi found in woodpecker nest-cavity trees or snags, which are predominated by fi lamentous Deuteromycetes fungi and yeast (Huss et al., 2002). There were only two species of white rot fungi, Coprinus sp. and Sporotrichum spp., and, as expected, none of brown rot fungi. The establishment of soft rot fungi may be infl uenced by the following factors. Firstly, the physical conditions inside the cavity may favor soft rot fungi. For instance, relative humidity inside the cavity tends to be higher than that of the outside (Poonswad, 1993), and the soft rot fungi tend to be found under high moisture conditions (Käärik, 1974). Secondly, after the pioneer species invades a wound of a tree, it normally causes heart and butt rot resulting in cavity formation. Then appropriate conditions in the cavity would facilitate the succession of other fungi. Thirdly, the succession of soft rot fungi appears to overwhelm the other groups of fungi. It could be that the cavities in this study were old (indicated by its large size, which accommodates a female hornbill and one chick or more), and that succession of the fungal community in the cavities has already occurred. Fourthly, either mutual or antagonistic interaction among fungi may affect the growth of one another on the host. Many soft rot fungi (e.g., Trichoderma spp., Penicillium spp., Gliocladium spp., Aspergilus spp.) established in this study were known to be antagonistic to the growth of Basidiomycetes (Ejechi, 1997, Huss et al., 2002). Therefore, the appearance of certain species of wood decay fungi is directly related to the time length and the decay rate.

Succession of fungi can take over the pioneer species through a series of organisms, which are capable of utilizing various components of wood as substrates over the course of time (Huss et al., 2002). From this information, it is clear that along the steps of wood degradation, different types of microorganisms are involved. Hence, rough dating of a cavity existence may be possible. Most of the fungi in this study which are known to have antagonistic activity to other fungi were predominantly found inside the cavities of Dipterocarpus and Cleistocalyx. This may be another reason the only one species of white rot fungi, Sporotrichum sp., was found in wood samples collected from inside nest cavities, whereas Coprinus sp. was only isolated from outside the cavity of Dipterocarpus in wet season. The establishment of Coprinus sp. in wet season can be interpreted as either this species is specifi c to Dipterocarpus tree or it is a seasonal occurrence. The positions where wood samples were collected from outside of the cavities of Dipterocarpus, Cleistocalyx, and Balakata in dry and wet seasons were relatively in the same region. This implies that a wound was fi rst created in dry season and then wood sample collections were repeated in wet season. Then, Coprinus, which was one of the pioneer species that created a cavity, may establish on Dipterocarpus tree during wet season.

Species diversity and host specifi city. – Two diversity indices adopted in this study in order to measure the diversity of fungal community on different host tree species weight on different factors. Simpson’s index is the highest for

fungal diversity of Dipterocarpus tree and second by that of Balakata (Table 2), while Shannon-Weiner index is the highest for fungal diversity of Balakata. Simpson’s index weighs towards equitability and is an intrinsic diversity index, giving the probability of any two individuals belonging to different species. Shannon-Wiener index is used in statistics as a diversity index. It is notoriously sample-size dependent and tends to be weighted slightly towards species richness. The result of this study corresponds to principle concept of these two indices. Balakata had the second highest diversity of fungi and its isolate number per species was evenly distributed, which was also indicated by the highest evenness value (Table 2). Whereas Cleistocalyx had markedly the lowest species diversity in dry season, and the number of isolates per species did not evenly distribute as it was indicated by the lowest evenness value (Table 2). In order to obtain accurate measurement of diversity, it is necessary to use more than one diversity index.

It is not possible to determine the host specifi city of fungi in this study due to inadequate samples of tree species studied, but we can present the exclusive species and wide host range species found under these limiting conditions. From the total of 68 fungal species found in this study, 35 species (51.5% of total) were exclusively found in all studied tree species (Appendix 1a). Among these fungi found from each studied tree, Cleistocalyx had the highest exclusive species (20 species or 29.4%, Appendix 1) indicating Cleistocalyx as a good host for many wood decay fungi. On the other hand, there were a few fungi (eight species or 11.8 % of total) that had wide host range (Dipterocarpus, Cleistocalyx and Balakata). About one third of the total species shared host trees of Dipterocarpus and Cleistocalyx (Appendix 1a). This may be one of the reasons these two tree species have similar numbers of cavities. Polishook (1996) suggested that much of host preference exhibited by micro-fungal decomposers may relate to physical and chemical characteristic factors rather than host specifi city by the sole taxonomic sense. Wood of some species have anti-fungal substances which resist certain species of fungi (Alfredsen et al., 2008) and this may result in different fungal community establishing in different wood.

On the other hand, May (1991) have reported that the high diversity in tropical forest ecosystems may better support those low host-specifi city fungi than those in low-diversity forests. It is further commented that the low density of individual host species in high-diversity forests limits the host preference or fungal population growth on suitable substrates (May, 1991). The exclusive fungi in Dipterocarpus and Cleistocalyx were found more in wet season. This clearly demonstrates the infl uence of the environmental factors rather than specifi c tree genera. From this study, it was inconclusive whether the exclusive fungi found on Dipterocarpus and Cleistocalyx were genuine host specifi c ones because they were from too small number of host tree species and from too few numbers of isolates (Appendix 1).

Overall, the total of 68 species found in this study tended to yield the maximum number of species richness of wood

105


decay fungi obtained from adequate number of host trees from each species (Fig. 3). To lengthen the species list, one should include more host tree species. Regardless of evenness, Cleistocalyx had greatest diversity of fungal species, whereas Balakata had the lowest. This substantial difference in establishment of fungi between Cleistocalyx and Balakata (Tables 2 and 3) tends to be affected by a number of combined factors, such as season and condition inside the cavity (e.g., temperature, aeration, humidity and substrate). The latter was not available in Balakata. Besides physical and biological (i.e., wood properties) factors, chemical compounds in wood may limit the growth of certain species. Wood of some species has anti-fungal substance, which resists certain species of fungi (Alfredsen et al., 2008) and results in different fungal community establishing in different wood.

ACKNOWLEDGEMENTS

We would like to express our sincere gratitude and appreciation to Dr. Suthep Wiyakrutta, Dr. Vithaya Meevootisom, for their creative guidance, discussions and constructive criticisms throughout this study. We also wish to sincerely thank Dr. Soraya Chaturongakul for helpful guidance; and Dr. Sumon Yudhasraparasithi for her suggestions and continuous discussion in statistical analysis.

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107

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hode

rma

ham

atum

0

0 0

0 0

2 2

0 4

0.33

So

ft r

ot

Tr

icho

derm

a ha

rzia

num

1

0 0

3 5

2 0

1 12

1

So

ft r

ot

Tr

icho

derm

a ko

ning

i 0

0 0

0 0

1 1

0 2

0.17

So

ft r

ot

Tr

icho

derm

a sp

iral

e 0

0 0

1 0

0 0

0 1

0.08

So

ft r

ot

Tr

icho

derm

a sp

1.

26

53

3 7

16

96

85

8 29

4 24

.52

So

ft r

ot

Ve

rtic

illi

um s

p.

3 4

0 0

0 9

5 0

21

1.75

Mic

roas

cace

ae

Soft

rot

Mic

roas

cus

sp.

0 0

0 0

0 0

3 0

3 0.

25

M

itosp

oric

Asc

omyc

ota

So

ft r

ot

G

liom

asti

x sp

. 0

1 0

0 0

1 0

0 2

0.17

So

ft r

ot

H

umic

ola

sp.

4 1

0 0

0 2

0 0

7 0.

58

Soft

rot

Pap

ulas

pora

sp.

0

0 0

0 0

1 1

0 2

0.17

So

ft r

ot

P

hom

a sp

. 0

0 0

0 0

0 2

0 2

0.17

108

Supa-Amornkul et al.: Wood decay in hornbill nestsA

ppen

dix

1. C

ont’

d

Cla

ssifi

cati

on

D

ry

W

et

To

tal

%P

hylu

m

Typ

e

G

ener

a/Sp

ecie

s D

ip

Cle

is

BB

D

D

DC

D

ip

Cle

is

BB

Is

o Is

o

of r

ot f

ungi

So

ft r

ot

Tr

icho

clad

ium

sp.

3

0 0

0 0

0 0

0 3

0.25

A

scom

ycot

a

M

itosp

oric

Cal

osph

aeri

acea

e

Soft

rot

Phi

alop

hora

ric

hard

siae

0

3 0

0 1

0 1

0 5

0.42

Mito

spor

ic D

erm

atea

ceae

So

ft r

ot

Tr

icho

spor

iell

a sp

. 0

4 1

0 0

0 0

0 5

0.42

Mito

spor

ic E

rem

omyc

etac

eae

So

ft r

ot

A

rthr

ogra

phis

cub

oide

a 0

0 0

0 0

0 0

2 2

0.17

Mito

spor

ic L

epto

shae

riac

eae

0 0

So

ft r

ot

E

pico

ccum

sp.

0

0 0

0 0

0 1

0 1

0.08

So

ft r

ot

C

onio

thyr

ium

sp.

0

0 0

0 0

4 0

0 4

0.33

Mito

spor

ic M

agna

port

hace

ae

Soft

rot

Phi

alop

hora

sp.

0

0 0

0 0

2 0

0 2

0.17

Mito

spor

ic M

icro

asca

ceae

So

ft r

ot

G

raph

ium

put

redi

nis

0 0

0 0

0 2

5 0

7 0.

58

Soft

rot

Gra

phiu

m s

p.

0 0

0 0

0 0

2 0

2 0.

17

M

itosp

oric

Pez

izac

eae

So

ft r

ot

O

edoc

epha

lum

sp.

0

0 0

0 0

0 1

0 1

0.08

Mito

spor

ic P

eziz

omyc

otin

a

Soft

rot

Toru

la h

erba

rum

0

0 0

0 0

0 1

0 1

0.08

So

ft r

ot

To

rula

sp.

3

0 0

0 0

0 0

0 3

0.25

Mito

spor

ic T

rich

ocom

acea

e

Soft

rot

Pae

cilo

myc

es v

icto

riae

0

0 0

0 1

0 0

7 8

0.67

So

ft r

ot

P

aeci

lom

yces

sp.

4

1 0

1 0

2 0

0 8

0.67

So

ft r

ot

P

enic

illi

um s

p.

10

4 2

2 0

0 1

0 19

1.

58

Soft

rot

Asp

ergi

llus

par

asit

icus

0

0 0

0 0

1 0

0 1

0.08

So

ft r

ot

A

sper

gill

us s

p.

17

2 2

0 0

1 7

4 33

2.

75

Soft

rot

Thy

sano

phor

a pe

nici

llio

ides

0

1 0

0 0

0 0

0 1

0.08

Nec

tric

eae

So

ft r

ot

C

ylin

droc

arpo

n oi

dium

0

0 0

0 0

1 1

0 2

0.17

So

ft r

ot

C

ylin

droc

arpo

n sp

. 3

1 0

1 0

5 12

0

22

1.83

So

ft r

ot

C

ylin

droc

ladi

um s

copa

rium

0

0 0

0 0

0 2

0 2

0.17

So

ft r

ot

F

usar

ium

ven

tric

osum

5

2 0

0 2

26

19

0 54

4.

5

Soft

rot

Fus

ariu

m m

onil

ifor

me

2 0

0 0

0 2

0 2

6 0.

5

Soft

rot

Nec

tria

sp.

0

0 0

0 0

0 3

0 3

0.25

Pleo

spor

acea

e

Soft

rot

Bip

olar

is o

ryza

e 0

0 2

0 0

0 0

8 10

0.

83

Sa

ccha

rom

ycet

acea

e

Non

rot

Can

dida

sp.

0

0 0

0 0

1 9

0 10

0.

83

Sc

opul

ario

psis

109


App

endi

x 1.

Con

t’d

Cla

ssifi

cati

on

D

ry

W

et

To

tal

%P

hylu

m

Typ

e

G

ener

a/Sp

ecie

s D

ip

Cle

is

BB

D

D

DC

D

ip

Cle

is

BB

Is

o Is

o

of r

ot f

ungi

So

ft r

ot

Sc

opul

ario

psis

asp

erul

a 0

0 0

0 0

0 3

0 3

0.25

Unc

lass

ifi ed

ord

er

Soft

rot

Mam

mar

ia s

p.

0 4

0 0

0 0

0 0

4 0.

33

Non

rot

Pha

eotr

icho

spae

ria

sp.

0 0

0 0

0 0

1 0

1 0.

08

X

ylar

iace

ae

0 0

So

ft r

ot

D

emot

opho

ra s

p.

0 1

0 0

0 0

0 1

2 0.

17

Bas

idio

myc

ota

Aga

rica

ceae

W

hite

rot

Cop

rinu

s sp

. 0

0 0

0 0

1 0

0 1

0.08

Mito

spor

ic A

gari

com

ycot

ina

W

hite

rot

Spor

otri

chum

sp.

3

7 0

0 0

40

9 0

59

4.92

O

omyc

ota

Pyth

iace

ae

Soft

rot

Pyt

hium

sp.

0

0 0

0 0

24

1 0

25

2.09

Z

ygom

ycot

a

C

unni

ngha

mel

lace

ae

Soft

rot

Cun

ning

ham

ella

sp.

0

0 0

0 0

10

0 0

10

0.83

Muc

orac

eae

0

So

ft r

ot

bs

idia

cli

ndro

spor

a 0

1 0

0 0

0 0

0 1

0.08

So

ft r

ot

A

bsid

ia s

p.

0 1

0 0

0 1

4 0

6 0.

5

Soft

rot

Gon

gron

ella

sp.

2

1 0

0 0

0 6

0 9

0.75

So

ft r

ot

M

ucor

sp.

0

2 0

2 0

2 3

0 9

0.75

Mor

tiere

lla

Soft

rot

Mor

tier

ella

sp.

0

0 3

0 0

1 6

0 10

0.

83

Tot

al i

sola

tes

136

195

21

22

37

429

318

41

1199

10

0

Tot

al s

peci

es

23

27

9 11

10

33

40

12

E

xclu

sive

No.

spe

cies

4

5 0

1 1

6 13

3

C

omm

on

D

ip +

Cle

is

10

18

D

ip +

BB

0

2

C

leis

+ B

B

1

1

Dip

+ C

leis

+ B

B

6

4

Dip

=D

ipte

roca

rpus

, Cle

is=

Cle

isto

caly

x, B

B=

Bal

akat

a, D

D=

dead

Dip

tero

carp

us, D

C=

dead

Cle

isoc

alyx

, Tot

al I

so=

tota

l is

olat

e, %

iso

=%

iso

late

.

110

Supa-Amornkul et al.: Wood decay in hornbill nestsA

ppen

dix

1.b)

Ide

ntifi

catio

n an

d nu

mbe

rs o

f fu

ngal

iso

late

s fr

om i

nsid

e an

d ou

tsid

e th

e ca

vitie

s of

Dip

terc

arpu

s an

d C

leis

toca

lyx.

Dry

W

et

Phy

lum

/Spe

cies

D

ipte

rocr

pus

Cle

isto

clay

x D

ipte

roca

rpus

C

leis

tocl

ayx

Insi

de

Out

side

In

side

O

utsi

de

Insi

de

Out

side

In

side

O

utsi

de

Asc

omyc

ota

B

ione

ctri

acea

e

G

lioc

ladi

um r

oseu

m

1

Can

dela

riac

eae

Can

dela

riel

la s

p.

1

1

Cap

nodi

ales

Fum

ago

sp.

3

C

haet

omia

ceae

Ach

aeto

miu

m s

p.

2

C

haet

omiu

m s

p.

1

Cha

etos

phae

riac

eae

Car

poli

gna

sp.

Cod

inae

a sp

.

2

Dip

odas

cace

ae

D

ipod

ascu

s sp

.

2

Gal

acto

myc

es g

eotr

ichu

m

1

Hyp

ocre

acea

e

A

crem

oniu

m s

p.

3

8

2

2 1

Cyl

indr

ocla

dium

sp.

2 3

4

5

1

F

usar

ium

cil

iatu

m

1

F

usar

ium

ros

eum

1

Fus

ariu

m s

olan

i 5

4

9

5 4

3

F

usar

ium

sp.

10

1

49

18

7

18

12

G

lioc

ladi

um v

iren

s 2

9

36

5

40

5

G

lioc

ladi

um v

irid

e

2

Gli

ocla

dium

sp.

22

2

24

1 89

11

21

1

Tric

hode

rma

ham

atum

2

2

Tric

hode

rma

harz

ianu

m

1

1 1

Tric

hode

rma

koni

ngi

1

1

Tr

icho

derm

a sp

iral

e

Tr

icho

derm

a sp

. 24

2

50

3 65

31

65

19

Vert

icil

lium

sp.

3

4

8

1 5

M

icro

asca

ceae

3

Mic

roas

cus

sp.

M

itosp

oric

Asc

omyc

ota

Gli

omas

tix

sp.

1

1

Hum

icol

a sp

. 4

1

1

1

P

apul

aspo

ra s

p.

1

1

P

hom

a sp

.

2

Tr

icho

clad

ium

sp.

3

111

THE RAFFLES BULLETIN OF ZOOLOGY 2011A

ppen

dix

1. C

ont’

d

Dry

W

et

Phy

lum

/Spe

cies

D

ipte

rocr

pus

Cle

isto

clay

x D

ipte

roca

rpus

C

leis

tocl

ayx

Insi

de

Out

side

In

side

O

utsi

de

Insi

de

Out

side

In

side

O

utsi

de

Asc

omyc

ota

M

itosp

oric

Cal

osph

aeri

acea

e

P

hial

opho

ra r

icha

rdsi

ae

3

1

Mito

spor

ic D

erm

atea

ceae

Tric

hosp

orie

lla

sp.

4

Mito

spor

ic E

rem

omyc

etac

eae

Art

hrog

raph

is c

uboi

dea

M

itosp

oric

Lep

tosh

aeri

acea

e

C

onio

thyr

ium

sp.

4

Epi

cocc

um s

p.

1

Mito

spor

ic M

agna

port

hace

ae

P

hial

opho

ra s

p.

2

Mito

spor

ic M

icro

asca

ceae

Gra

phiu

m p

utre

dini

s

2

5

Gra

phiu

m s

p.

1 1

M

itosp

oric

Pez

izac

eae

Oed

ocep

halu

m s

p.

1

Mito

spor

ic P

eziz

omyc

otin

a

To

rula

her

baru

m

1

Toru

la s

p.

3

Mito

spor

ic T

rich

ocom

acea

e

A

sper

gill

us p

aras

itic

us

1

Asp

ergi

llus

sp.

15

2

2

1

7

P

aeci

lom

yces

vic

tori

ae

P

aeci

lom

yces

sp.

4

1

2

Pen

icil

lium

sp.

7

3 4

1

Thy

sano

phor

a pe

nici

llio

ides

1

N

ectr

icea

e

C

ylin

droc

arpo

n oi

dium

1

1

Cyl

indr

ocar

pon

sp.

3

1

5

10

2

C

ylin

droc

ladi

um s

copa

rium

2

Fus

ariu

m m

onil

ifor

me

2

2

F

usar

ium

ven

tric

osum

5

2

17

9

8 11

Nec

tria

sp.

3

Pl

eosp

orac

eae

Bip

olar

is o

ryza

e

Sacc

haro

myc

etac

eae

Can

dida

sp.

1

9

Sc

opul

ario

psis

Scop

ular

iops

is a

sper

ula

3

112


App

endi

x 1.

Con

t’d

Dry

W

et

Phy

lum

/Spe

cies

D

ipte

rocr

pus

Cle

isto

clay

x D

ipte

roca

rpus

C

leis

tocl

ayx

Insi

de

Out

side

In

side

O

utsi

de

Insi

de

Out

side

In

side

O

utsi

de

U

ncla

ssifi

ed o

rder

Mam

mar

ia s

p.

3 1

Pha

eotr

icho

spae

ria

sp.

1

Xyl

aria

ceae

Dem

otop

hora

sp.

1

Bas

idio

myc

ota

A

gari

cace

ae

C

opri

nus

sp.

1

M

itosp

oric

Aga

rico

myc

otin

a

Sp

orot

rich

um s

p.

3

7

26

14

9O

omyc

etes

Py

thia

ceae

Pyt

hium

sp.

24

1Z

ygom

ycot

a

Cun

ning

ham

ella

ceae

Cun

ning

ham

ella

sp.

10

M

ortie

rella

ceae

Mor

tier

ella

sp.

1

6

M

ucor

acea

e

A

bisi

dia

clin

dros

pora

1

Abi

sidi

a sp

.

1

1

4

Gon

gron

ella

sp.

2 1

6

Muc

or s

p.

2

2

2 1

Tot

al i

sola

tes

122

14

190

5 33

8 91

25

5 63

Tot

al s

peci

es

21

7 26

3

29

15

37

14

113


Appendix 2. Properties of wood sampled from Dipterocarpus gracilis, Cleistocalyx nervosum and Balakata baccata (referred to as D. costata, Syzygium cumini, Sapium baccatum, irrespectively in Thonanont et. al, 1985).

Tree species Moisture content Toughness Strength Hardness % kg-m Kg/cm2 kg

Moderate hardwood with high strength

D. gracilis 12 4.03 661 772

Moderate hardwood

C. nervosum 12 2.86 473 809

Softwood

B. baccata 187 1.90 167 176

wood decay fungi in hornbill nest cavities in khao yai...

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